Biochemical composition of the Atlantic bonito Sarda sarda from the Aegean Sea (eastern Mediterranean Sea) in different stages of sexual maturity

The content (% wet mass) in water, ash, lipid, crude protein, DNA and RNA of different tissues was determined during sexual maturation of bonitos Sarda sarda from the Aegean Sea. A total of 220 specimens were collected in the following stages of sexual maturity: immature, resting, developing, mature, spawning and spent. Highest lipid levels in the white muscle, red muscle and liver were measured in immature specimens, while lowest levels were found in spawning bonitos. The gradual percentage of lipid reduction from immature to spawning bonitos was relatively higher in the liver (females 71·2% and males 64·4%) than in the white (females 59·2% and males 53·5%) and red (females 62·1% and males 51·7%) muscle. Lipid levels in the gonads increased gradually from the immature to spawning stage. The decrease of lipid in the somatic tissues was more intense in females than in males, and gonadal lipid content was higher in females than in males. There was a strong reverse correlation between water and lipid percentage in all tissues. Protein content decreased significantly only in spawning bonitos. The percentage of protein reduction from immature to spawning stage was relatively higher in males than in females in both white (females 3·4% and males 4·6%) and red (females 4·6% and males 5·1%) muscles. Protein content in the liver was significantly lower than in the other tissues, being highest in mature females. Gonadal protein content in females increased with maturation and decreased after spawning. The content in ash exhibited considerable stability. The RNA:DNA ratio exhibited a similar pattern of variation in both muscles. The RNA:DNA ratio increased during gonadal development gradually from the developing to spent stage. It was concluded that in S. sarda during gonadal development, there was an increase in gonadal lipid accompanied by a decrease in somatic tissue lipid reserves. Thus, reproductive inactive bonitos have more lipid in their edible part and a higher nutritional value than active ones.     

Reproductive energy expenditure and changes in body morphology for a population of Chinook salmon Oncorhynchus tshawytscha with a long distance migration

Energetic demands of a long freshwater migration, extended holding period, gamete development and spawning were evaluated for a population of stream‐type Chinook salmon Oncorhynchus tshawytscha. Female and male somatic mass decreased by 24 and 21%, respectively, during migration and by an additional 18 and 12% during holding. Between freshwater entry and death after spawning, females allocated 14% of initial somatic energy towards gonad development and 78% for metabolism (46, 25 and 7% during migration, holding and spawning, respectively). Males used only 2% of initial somatic energy for gonad development and 80% on metabolic costs, as well as an increase in snout length (41, 28 and 11% during migration, holding and spawning, respectively). Individually marked O. tshawytscha took between 27 and 53 days to migrate 920 km. Those with slower travel times through the dammed section of the migration corridor arrived at spawning grounds with less muscle energy than faster migrants. Although energy depletion did not appear to be the proximate cause of death in most pre‐spawn mortalities, average final post‐spawning somatic energy densities were low at 3·6 kJ g^?1 in females and 4·1 kJ g^?1 in males, consistent with the concept of a minimum energy threshold required to sustain life in semelparous salmonids.     

Reproductive cycle and reproductive output of the sea urchin Loxechinus albus (Echinodermata: Echinoidea) from Beagle Channel, Tierra del Fuego, Argentina

Reproductive cycle, frequency and duration of spawning, energetic content of gonads, and reproductive output of the common green sea urchin Loxechinus albus were analyzed in the Beagle Channel (Tierra del Fuego) between May 2004 and May 2005. Gonad indices (GI, percentages of gonad mass in total body mass) were significantly higher in March, April, July, and August than in November and May, thus showing a negative correlation with the photoperiod. Highest GI values of mature individuals were observed in August, and spawning occurred from September to December. In females, the mass-specific energy content of gonads (ECG) was highest in spawned gonads and lowest in mature ones, while in males ECG values were higher in immature stage and lower in premature and mature stages. High ECG values can be explained by the abundance of nutritive phagocytes. Both ECG and total gonad energy content (TECG) were higher in females than in males. Mean reproductive output was 7.28% for females and 6.15% for males (expressed as the difference between mean GI of mature and spawned gonads) and 25.02 kJ for females and 19.26 kJ for males (expressed as the difference between mean TECG of mature and spawned gonads).     

Reproduction of the lambari Astyanax scabripinnis (Jenyns, 1842) (Pisces: Characidae) in a small stream in Southeastern Brazil

The reproduction of Astyanax scabripinnis captured bimonthly in the Cristais small stream (20^o00'03'S and 43^o56'46'W), in the Das Velhas River basin, was studied using gross anatomical and histological techniques. Standard length of females (56 ± 8 mm) was longer than males (49 ± 6 mm). The diameter of the nucleus of the primary spermatogonia was 7.5 ± 0.9  μ m and of the spermatozoon head 1.8 ± 0.4  μ m. For the females, the oogonia nuclei measured 10.2 ± 2.2  μ m and the vitellogenic oocyte nuclei measured 675.0 ± 66.7  μ m. The following stages of the reproductive cycle were established: developing, mature, partially spent/spawned and completely spent/spawned. Maximum number of males (61.3%) and females (58.2%) at the mature stage was observed during November/December. The gonadosomatic index of males and females was highest during the mature stage (2.4 ± 0.6; 7.5 ± 1.3, respectively) and decreased in subsequent stages. In females, the hepatosomatic index was highest during the developing stage (2.8 ± 0.3), and slightly decreased in the following stages. The highest stomach repletion index found for males was observed during the partially spent stage (2.0 ± 1.2), while for the females, this index varied discretely between the reproductive stages. The highest values of Fulton's condition factor were registered during the mature stage in both sexes (2.9 ± 0.4 for males; 3.0 ± 0.3 for females). The continuous reproductive period and the presence of partially spawned females during the entire sampling period indicated that the species has fractional spawning.     

Energetic cost of spawning in male and female Atlantic salmon (Salmo salar L.)

Energetic expenditure during spawning of male and female 1 -sea-winter Atlantic salmon, Salmo salar L., was measured. Before spawning, the somatic energy content per unit weight did not differ between the sexes. The testes content offat was 0.24, of carbohydrate 8.89, of water 1.21 and of ash 1.61 times as high as that of the ovaries. Just prior to spawning, mean gonadosomatic index (GSI; wet weight, %) in males was 4.36 and in females 20.26, and expressed as energy ratios (kJ, %) 4.47 and 27.75, respectively. During spawning the energy loss of male soma was higher (35.57%) than that of females (25.00%). This was a result of higher loss of fat in males than in females. Total energy cost of spawning in males and females was on average 51.8 and 51.4%, respectively.     

Reproductive biology of albacore Thunnus alalunga

Reproductive variables in albacore Thunnus alalunga were evaluated by gonad histology in samples of 132 males (58–118 cm fork length, L_F) and 112 females (59–101 cm L_F) that were collected from the western North Pacific Ocean from 2001 to 2006. In the sex ratio examination, males greatly outnumbered females in large adult fish (L_F > 100 cm). Thunnus alalunga exhibited a protracted spawning period from March to September in the waters off eastern Taiwan and the Philippines, and the peak spawning activity occurred in March and April. Minimum sizes associated with the classification of mature fish were 78 and 83 cm L_F for males and females, respectively. In addition, the largest L_F of immature fish were 93 cm for males and 94 cm for females. The spawning frequency estimate in April was 1·7 days. Batch‐fecundity estimates of 21 females (89–99 cm L_F) ranged between 0·17 and 1·66 million eggs (mean ±s.d . = 0·94 ± 0·43). The relative fecundity estimates of the 21 females ranged between 9·2 and 92·4 oocytes g^?1 body mass (mean ±s.d . = 50·5 ± 22·8). The results presented in this study provide increased information regarding this species' reproductive‐related characteristics than are currently available in stock status determinations.     

社群等级对金乌贼行为表型及能量代谢的影响

采用实验生态学方法,在室内水槽条件下研究了金乌贼(Sepia esculenta Hoyle,1885)繁殖过程中社群等级的形成对其行为表型和能量代谢的影响,分析测定了不同优势等级雌雄个体腕部肌肉和性腺组织中己糖激酶(Hexokinase, HK)、丙酮酸激酶(Pyruvate kinase, PK)、乳酸脱氢酶(Lactate dehydrogenase, LDH)、苹果酸脱氢酶(Malate dehydrogenase, MDH)、柠檬酸合酶(Citrate synthetase, CS)活性以及乳酸(Lactic acid, LD)含量。结果显示:(1)金乌贼繁殖期不同优势等级雌雄个体之间行为表型具有显著差异,优势雄性个体游动悬浮、争斗时间显著高于劣势个体,而优势雌性个体静止伏底时间高于劣势个体,游动悬浮时间低于劣势雌性;(2)优势雄性个体在争斗过程中主要通过无氧代谢提供能量,而处于游动悬浮状态时通过有氧代谢提供能量。主要表现在优势雄性个体肌肉中无氧代谢酶(PK、HK、LDH)活性显著高于劣势个体(P<0.05),有氧代谢酶(MDH、CS)活性也显著高于劣势个体,雌性个体之...     

Biochemical content, energy composition and reproductive effort in the broadcasting sea star Asterias vulgaris over the spawning period

To assess sex differences in reproductive effort, we examined the biochemical composition and energetic content of the principal body components of the broadcast spawning sea star Asterias vulgaris in the Mingan Islands in the northern Gulf of St. Lawrence, eastern Canada. The body wall was the most stable body component, showing no variations in mass or in lipid and protein content (and total energetic content) between sexes or during spawning. Patterns in the gonads differed between sexes and with spawning. The lipid, protein and carbohydrate content of the ovary dropped during spawning, while only the protein content of the testis decreased significantly. Reproductive effort, expressed as loss of energy in the gonads during spawning for an individual weighing 10 g in underwater mass (8.2 cm in radius), was six times greater in females (49.5 kJ) than males (7.9 kJ). The energetic content of the pyloric caeca also decreased during spawning, by 17.7 kJ in females and 21.5 kJ in males, mainly due to a decrease in lipids. If this decrease is included as reproductive effort, it lessens the gender difference. The caecum decrease possibly represented expenditures due to formation of aggregations or the expulsion of gametes during spawning. Effectively, we observed aggregations during a massive spawning in this population. The sex ratio did not differ from 1:1 in all size classes sampled. This suggests that, unless males suffer higher mortality, females manage to allocate as much energy to somatic growth as males, possibly by feeding at higher rates to compensate for their higher reproductive effort. Stomach protein content tended to be higher in females than males and may indicate greater muscular development to facilitate digestion.     

Successful gonadal development and maturation of tench (Tinca tinca L.) in small concrete ponds

The experiments were performed in a tench farm from autumn until the spawning season (June–July). Tench broodstocks from natural habitats were maintained in 25 × 6 × 1 m concrete ponds and fed on commercial trout pellets. Females and males were separated and maintained under natural photoperiod and temperature conditions at densities around 2 kg m^?2. Water flow throughout was supplied at the rate of 15 L s^?1. When females showed external signs of advanced gonadal development, induction of spawning was made by luteinizing hormone releasing factor (LH‐RH) synthetic analogue administration at three different periods of the reproductive season (June–July). A single intramuscular injection (20 μg kg^?1 body weight) was administered to 110 mature females selected from a total of 150. The females were stripped 42 h (22°C) after hormone administration. The mean rate of stripped females to the number injected was 77%. Mean relative egg weight in relation to the weight of the stripped females was 5.61%. More than 90% of the males provided semen without hormonal induction. Differences in egg production and external egg quality were observed at different times of the spawning period. It was proven that tench maintained in small concrete tanks and fed on artificial diets were able to reach gonadal maturation.     

Effects of varying temperature and food availability on growth and reproduction in first‐time spawning female Atlantic cod

The somatic growth, sexual maturation and fecundity of individually marked first‐time spawning female Atlantic cod Gadus morhua were examined under different varying temperature and feeding regimes over the months preceding spawning. A negative correlation between somatic and oocyte growth was found, reflecting the changing energy allocation pattern. Nevertheless, the somatic growth of mature individuals was at least as high as those of immature fish over the period of vitellogenesis. Potential fecundity was positively correlated with body size, but neither temperature or feeding regime significantly affected this relationship. Consequently, fish with unlimited feeding opportunity invested more energy into somatic growth during vitellogenesis compared to those held under a restricted ration. This indicated that once Atlantic cod had made the decision to invest in first reproduction, they allocated a certain amount of energy relative to their size into egg production and any surplus was invested into somatic growth. Low temperature led to an arrest in the onset of vitellogenesis and significantly affected the number of females that matured.     

The seasonal changes in somatic energy content of Gulf of Alaska yellowfin sole, Pleuronectes asper

Seasonal measurement of body energy content was made for Pleuronectes asper (Pallas, 1814) from the Gulf of Alaska. Whole body energy content of complete fish (～ 3472 J g^?1 wet wt) was minimal in May for females as their overwintering phase ended, then increased to ～ 4456 J g^?1 prior to spawning. The ovarian index [(g.w./t.w.) × 100] and energy content of ovaries (J g^?1) was highest in June and May respectively, and then declined markedly by August as spawning occurred. Throughout the year male whole body energy content of complete fish ranged from 3351 to 4590 J g^?1 with the lowest values in May and highest values occurring during June to September, the feeding season. The testes index [(g.w./t.w.) × 100] and total energy content of testes (J g^?1) were high in March and lowest during June and July. On a weight-specific basis, males and females had similar whole body energy values throughout the year. Juveniles followed the same seasonal trends in energy storage as adults and had similar whole body energy values. Whole body energy content was linearly related to wet and dry weight condition factor with r² values of 0.70 and 0.87, respectively. Dry body weight as percent of wet body weight was the best predictor of body energy (r²=0.91). Yellowfin sole had an annual energy cycle with energy accumulation and growth from May to September. Thereafter they utilized stored energy for metabolic and reproductive needs. Spawning began in late May or early June and fish were spent by August. Whole body energy content increased by 28, 33 and 35% between May and June, for females, juveniles, and males, respectively, the most dramatic change during the year long survey. This suggests that intense feeding in May must be an important aspect of their energy storage cycle.     

Competitive mate searching in male common toads, Bufo bufo

The reproductive behaviour of a population of individually marked toads Bufo bufo was studied at a pond where males outnumbered females by between four and five to one. There was intense competition between the males for mates and only 20·5 % of them bred successfully. Of the successful males, 38·5 % got mates by fighting and displacing other males from the backs of females (takeovers). Larger males enjoyed greater reproductive success because they were stronger and better able to achieve takeovers. When competing for females, some males searched at the spawn site while others searched away from spawn. The numbers searching in the two areas can be predicted by a model which assumes that unpaired males distribute themselves so that there is a spatial ESS, where individuals have equal expectations of finding a female both at and away from the spawning ground.     

Relation between age–sex classes and dietary selection of wild Japanese monkeys

Dietary selection by wild Japanese monkeys (Macaca fuscata yakui Kuroda) was examined in relation to body size. The monkeys were classified into three age–sex categories: adult males, adult females and immatures excluding suckling infants. Time spent feeding did not differ between age–sex classes, although time spent moving was longer in lighter classes. Heavier individuals fed relatively more on mature leaves, while lighter individuals fed on insects more frequently. Mature leaves were more abundant but had lower energy content than other food items in the forest. Heavier monkeys seemed to feed on greater amounts of lower quality food in this species. This finding agrees with the Jarman–Bell principle on ungulates.     

Differences in δ13C and δ15N stable isotopes in the pearly razorfish Xyrichtys novacula related to the sex,location and spawning period

In the present study, Xyrichtys novacula (Labridae) were sampled at five locations around the islands of Ibiza and Formentera (western Mediterranean Sea). Isotopic signatures of δ¹³C, δ¹⁵N and the C:N ratio were analysed in relation to locality, sex and size differences. δ¹³C and δ¹⁵N partitioning was also studied in the reproductive spawning period. There were significant differences in the δ¹³C signature between localities for both sexes, but not for δ¹⁵N. Sex differences were also found with a mean ±s.e . value of ?17·38 ± 0·06‰δ¹³C and 8·36 ± 0·05‰δ¹⁵N for females and ?17·17 ± 0·07‰δ¹³C and 8·80 ± 0·06‰δ¹⁵N for males. Increasing total length in both sexes was positively correlated with δ¹⁵N enrichment and a significant positive linear regression was established for both variables. During the reproductive spawning period, there were changes in δ¹³C fractioning with enrichment in postspawning females and males (with respect to prespawning and spawning periods) and δ¹⁵N impoverishment in postspawning females (with respect to prespawning and spawning periods). Xyrichtys novacula uses local food sources, as confirmed by δ¹³C and δ¹⁵N, and females and males use different food sources, thus avoiding intraspecific competition. This was confirmed by δ¹⁵N enrichment as size increased. Spawning leads to special requirements for gonad maturation, which is reflected in the isotopic signatures for both sexes.     

大鳍鳠鱼体能量密度及其预测模型

于2000年11月至2001年10月由嘉陵江收集野生大鳍鳠成鱼100尾,其中冬季(12月)、春季(3月)、繁殖前(6月)、繁殖后(8月)、秋季(10月)雌雄各10尾。测定了这10个样本的鱼体能量密度及生化组成。统计分析结果表明：鱼体各生化成分含量,能量密度的性别间差异在各个季度均不显。在雌性群体中,秋季样本的蛋白质含量(16．23％),脂肪含量(5．84％),干物质含量(26．67％)和能量密度(6．14kJ／g)显高于其余各样本的相应指标。春季样本中鱼体脂肪含量和能量密度最低(2．39％,4．55kJ／g)。在雄性群体中,秋季样本的蛋白质含量(16．34％)与能量密度(5．85kJ／g)最高;春季样本中脂肪含量(1．3l％)、干物质含量(22．49％)、能量密度(4．16k．J／g)最低。各样本能量密度均分别与干物质和脂肪含量呈显的直线相关关系。通过协方差分析,将雌性群体中冬季、春季、秋季以及繁殖后样本共4个样本的能量密度与干物质含量的公共回归方程作为以干物质含量(D)预测大鳍鳠雌性群体在非繁殖期能量密度(E)的预测模型：E=-5．573 0．437D;将繁殖前雌性样本的能量密度与干物质含量的回归方程作为预测大鳍鳠雌性群体在繁殖期的能量密度的预测模型：E=-0．605 0．250D;将冬季、春季、秋季以及繁殖后雄性4个样本的能量密度与干物质含量的公共回归方程作为预测大鳍鳠雄性群体在非繁殖期的能量密度的预测模型：E=-6．046 0．456D;将繁殖前雄性样本的能量密度与干物质含量回归方程作为预测大鳍鳠雄性群体在繁殖期的能量密度的预测模型：E=-3．64 0．366D。     

Egg production of the marine,planktonic copepod,Calanus pacificus Brodsky: Laboratory observations

When acclimated to a continuous, superabundant food supply and constant temperature, Calanus pacificus Brodsky females produce eggs at a weight-specific rate ranging from 0.13 · day^?1 at 8°C to 0.21 · day^?1 at 15°C. Maximum weight-specific egg production rates do not change with seasonal changes in female body size. The relationship between egg production rate and food concentration is hyperbolic, with threshold and critical concentrations that are high relative to other species for which data are available. Food concentration and temperature influence spawning frequency (i.e., the time required for oocytes to mature) much more than the number of eggs in a single spawning event (i.e., clutch size). Clutch size is significantly related to female body size.     

Seasonal changes in energy and the energy cost of spawning in Gulf of Alaska Pacific cod

In the Gulf of Alaska, adult Pacific cod exhibited an annual cycle of condition, gonad index and liver index in which maximum values occurred in ripe fish in March and minima in July. About 30–31 % of prespawning stored energy was expended during the spawning effort. The energy associated with spawning derived from liver (24% and 18%), somatic tissue (22% and 33%) and gonad (53% and 48%) for females and males, respectively. Liver index and gonad index at the time of sampling were directly related in females, but in males gonad index was best related to liver index 1–3 months earlier.
The Pacific cod is very similar to the Atlantic cod in terms of energy cycling, maximum gonad sizes, energy expended during spawning and gonadal contribution to energy expenditure. However, in Pacific cod, somatic tissue contributes markedly to energy expended during reproduction. The Pacific cod cod differs from the walleye pollock with respect to gonad index (13% and 20%ν. 20% and 8% for females and males, respectively), spawning weight loss (25%ν. 38%), liver energy loss during spawning (71%ν. 55%) and energy cost of spawning.     

Energy allocation for reproduction in a marsupial arboreal folivore,the common ringtail possum (Pseudocheirus peregrinus)

This study examines the annual energetics of a small folivorous marsupial, Pseudocheirus peregrinus. Particular attention was given to the energy and time allocated to reproduction by the females. Daily energy expenditure was measured directly using the doubly labelled water technique. Energy transferred to the young via the milk was estimated from information on milk composition and production. There was no significant seasonal variation in the energy expenditure or water influx of males or females. The mean daily energy expenditure of a 1-kg non-lactating adult ringtail possum was 615 kJ day^–1 or 2.2 times standard metabolic rate. Females showed significant changes in daily energy expenditure according to their reproductive status. Without the burden of lactation the total annual energy expenditure of an adult female was estimated as 212.4 MJ kg^–1 year^–1. The total annual energy expenditure of a female rearing two young was 247.5 MJ kg^–1 year^–1, with the late stage of lactation constituting the most energetically expensive period accounting for 30% of the total yearly energy expenditure during 24% of the time. Total metabolisable energy allocation during reproduction (22 MJ kg) was similar to estimates available for other herbivores, although, the peak metabolisable energy allocation during lactation (759 kJ day^–1) was lower than values available for other herbivores. The total energy requirement for reproduction (metabolisable energy plus potential energy exported to young via milk) suggests that the ringtail possum also has a relatively low overall energy investment in reproduction. It is suggested that the lactational strategy of the ringtail possum has been selected in order to spread the energy demands of reproduction over time due to constraints on the rate of energy intake imposed by a leaf diet and/or to prolong the mother-young bond. The strategies a female ringtail possum may employ to achieve energy balance when faced with the energy demands of reproduction are discussed.     

Estimation of food intake and ingested energy in Daubenton’s bats (Myotis daubentonii) during pregnancy and spermatogenesis

We studied food intake of and estimated ingested energy in female and male Myotis daubentonii during the periods of pregnancy (period 1, 8 May–4 June) and of intense spermatogenetic activity (period 2, 24 July–22 August) over 8 years (1996–2003) in central Germany. We used radiotelemetry to determine the time spent foraging and marked animals with chemiluminescent light-sticks to determine prey attack rates. Body length, body mass, moisture content, and caloric content of chironomids, the main prey of Daubenton’s bats, were measured to estimate the nightly food intake and, in consequence, energy intake. Pregnant females spent significantly more time foraging than males during period 1 and females during the post-lactation period. In contrast, male foraged longer during the period of highest spermatogenetic activity than during late spring and also significantly longer than post-lactating females. Based on a mean number of 8.3 prey attacks per minute, the time spent foraging, and a capture success rate of either 50 or 92%, calculated intake values with a feeding rate of 7.6 insects per minute (=92% capture success) were more consistent with literature data for other insectivorous bats than that of values calculated on the basis of a capture success rate of 50%. In the high capture-success model, calculated insect intake of female bats was 8.0 g during pregnancy and 4.9 g per day during post-lactation, providing 5.0 and 3.0 kJ of ingested energy per gram body mass per day. Calculated intake of male bats was 3.6 g insects per day during late spring and 8.0 g during period of intensive spermatogenesis, providing 2.6 and 5.7 kJ of ingested energy per gram body mass.     

Migratory Atlantic salmon as vectors for the transfer of energy and nutrients between freshwater and marine environments

SUMMARY 1. Annual energy, carbon, nitrogen and phosphorus fluxes across the river mouth by Atlantic salmon were estimated for 18 years (1976–94) in the Norwegian River Imsa. The total energy content of the emigrating smolts in each year varied considerably with a mean value of 237 × 10³ kJ. That of returning adults also varied between years with a mean value of 141 × 10⁴ kJ. One‐sea‐winter salmon (grilse) made up 65% of the total energy content of the spawners in the river. Dead carcasses remaining in the river after spawning were estimated to have a mean annual energy content of 175 × 10³ kJ. 2. The net annual energy flux from the sea to the river varied between 48 × 10³ kJ (1987) and 152 × 10⁴ kJ (1989) with a mean of 616 × 10³ kJ, and a coefficient of variation of 67%. Average net marine import of the returning adults was 83 × 10⁴ kJ year^?1 with a coefficient of variation of 52%. Mean annual export of C, N and P to sea by the smolts was 595, 131 and 22 kg, and by kelts 1535, 352 and 70 kg, respectively, whereas gross import via the adults was 3176 kg C, 735 kg N and 132 kg P. The annual flux across the river mouth was 1046 kg C, 253 kg N and 39 kg P. The net marine import were 1585 kg C, 371 kg N and 60 kg P. The net flux was estimated at 0.2% for nitrogen and 5% for phosphorus of the total river load. 3. The energy flux caused by Atlantic salmon spawning in the River Imsa was relatively high because the general nutrient load in the river is low. Thus, even though most Atlantic salmon survive spawning, their contribution to the nutrient flux in the river is significant.